rieann
A3
)tection
January 2001
•
MONITORING PONDEROSA PINE DECLINE AND MORTALITY
ON THE FLATHEAD INDIAN RESERVATION
Establishment Report and 5-Year Results
•
Blakey Lockman and Tom Corse
INTRODUCTION
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Mortality in ponderosa pine from annosum root
disease (Heterobasidion annosum (Fr.) Bref) was
first diagnosed on the Flathead Indian
Reservation (IR) in the mid 1970's (Williams and
Haglund 1976; see Appendix for biology of
annosum root disease). Extensive root disease
and/or drought symptoms and associated
mortality were recognized in the Dry Fork area
and nearby Sunrise area on the Reservation's
west side in the late 1980's, and appeared to
increase significantly in 1990. Mature pines
scattered throughout that area had been dying at
an unknown rate. Many crowns were thin, with
needle retention lower than expected; terminal
growth was poor; and in some cases, foliage
color was chlorotic or appeared to have a silver
cast. Annosum root disease had been identified
as a causal agent in some of these situations.
In 1993, we established permanent plots to
monitor the decline of ponderosa pine on the
west side of the Flathead IR. The main
objectives of the study were: (1) determine rate
of decline of individual trees; (2) identify
symptoms which can be used to predict time of
mortality; and (3) determine mortality rate over
time. All of these objectives will help the Tribes
United States
Department of
Agriculture
Forest
Service
Northern
Region
manage their ponderosa pine. Large ponderosa
pine have great significance to tribal members,
but economics dictate harvesting of trees before
they die to optimize monetary value. We also
hoped to determine the overall cause of decline
and mortality.
METHODS
Plot Selection
We investigated the possibility of using annual
aerial videography to monitor mortality with the
Methods Application Group (MAG), Fort
Collins (now called Forest Health Technology
Enterprise Team, Fort Collins). We determined
aerial video technology was too much in its
infancy to use as the main survey tool. MAG
considered using this project as a test site for the
usefulness of aerial videography to monitor
annual mortality, so we attempted to design
ground plots to compliment a future aerial video
test project.
We created a random number list using UTM
coordinates from the Ill's west side under 4,400
feet elevation (R24W and T19N north to T24N).
This general area was selected by Tom Corse, IR
forester, as the area of most concern.
200 East Broadway
P.O. Box 7669
Missoula, MT 59807
with directions for locating them. Transects were
established in 1993.
We selected the first 23 UTM coordinates from
the random number list and then delineated 1/4mile wide flight lines on 1:24,000 topographic
maps. Each flight line was drawn from the
selected UTM coordinate north to the next UTM
coordinate. The first 18 flight lines that were
entirely on IR land, were forested, and mostly
under 4,400 feet elevation were then selected for
aerial photography.
At the time of plot establishment, the Tribes
requested we include a transect in an area of
known annosum infection. They selected the site
and we verified the presence of annosum fruiting
bodies on the roots of overstory trees and
seedlings/saplings. The Tribes located this nonrandomly selected transect (Annosum transect)
adjacent to the leading edge of an active H.
annosum root disease center. This transect was
not flown and was analysed separately.
In June 1993, these 18 flight lines were flown
and 1:6000 aerial photography was taken. Three
flight lines were discarded because they were in
an active timber sale (Welcome Springs), which
left 15 flight lines.
Data Collected
At plot establishment, all trees 15 inches or
greater in diameter at breast height (d.b.h.)
within boundaries were tagged at the base with
an aluminum numbered tag. Their general
location was also noted on a transect grid map.
Crowns of all trees were given a numerical rating
for needle complement, needle length, needle
color, and twig and branch condition. The
highest cumulative rating a crown could receive
was 9 (Table 1). A high rating indicated a tree in
poor condition. This rating system was based on
risk rating systems developed for predicting
mortality of ponderosa pine from bark beetles
(Salman and Bongberg 1942) and oxidant air
pollution (Miller 1973).
The photo which best represented each flight line
was selected. A scaled overlay which represented
an 8-acre transect 264 feet wide (east/west) by
1,320 feet long (north/south) was placed over
each photo to delineate ground sampling.
Starting point for each transect was the south end
of overlay. Each photo was then pinpricked and
the spot was located on the ground. Ground
transects ran due north for 1,320 feet from
starting points. Rebar was placed at starting and
ending points on all transects, and reference trees
were painted and tagged with an aluminum tag at
approximately 4.5 feet above ground. Starting
points were located on maps and photos, along
2
Table 1. Ponderosa pine crown rating system.
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SYMPTOMS
_
1"- RATING
e oini3 em6n
a. normal (3+ years retention)
b. 2-year needle retention
c. 1-year needle retention (bottle brush))
0
1
2
a. normal
b. shorter than normal throughout crown
c. needles shorter in upper crown, normal below a definite contrast
.
6 e
a. normal
b. off color (moderately chlorotic)
c. fading, more yellow than green
..
an
a. no twigs or branches dead
b. a few scattered dead or dying branches or twigs in crown, with <10% occurring in uper V2 of crown
c. 10-20% of twigs and branches in upper of crown dead
d. greater than 20% of twigs and branches inupper 'A of crown dead
3'
4N;IO..,*14:g
COMU- LAT'CROWNATIN
8-'
The following information was also collected
and recorded for each tree: d.b.h., crown ratio,
crown class, and damages using Region 1
Timber Stand Exam damage codes (Anonymous
1985: 415.4-415.51). A standard stand exam plot
(Anonymous 1985) was then nested on the
transect center line at three locations: 264, 660,
and 1,056 feet from each starting point. A
variable-radius plot using 10 BAF (basal area
factor) was used to tally trees 5 inches d.b.h. and
larger, and a nested 1/100-acre fixed plot was
used to count regeneration and estimate ground
coverages.
-
0
1
2
0
1
2
0
1
2
3
9
as close to ground level as possible and
incubated in a moisture chamber to verify the
presence of the asexual stage of the fungus,
Spiniger meineckellus (A.J. Olson) Stalpers.
Transects were revisited each year, except 1997,
when time and financial constraints prevented a
remeasurement. Mortality was recorded at these
annual visits, as well as any significant changes.
In 1998, in addition to mortality, crowns of all
live tagged trees were rated again and damages
were again recorded. Any missing tags were
replaced, and reference trees were repainted as
necessary.
In addition to the above data, an informal survey
was done on each transect looking for the
presence of H. annosum. Cut stumps were
investigated for typical decay and/or fruiting
bodies, and seedling and sapling mortality was
checked for presence of button conks. When
possible, mortality trees were sampled to
determine the presence of H. annosum. Dead
trees were cut and 2-inch thick discs were taken
Statistical Analysis
One-way analysis of variance was used to test
for significant differences between live and dead
trees. T-tests for paired samples were used to test
for significant differences between 1993
measurements and 1998 measurements of live
trees (SPSS 1994).
3
Table 2. Transect characteristics at time of plot establishment.
•
# Ponderosa
Elevation
(feet
' 0071*,
3600
Transect
#
2
Slope
.240;T;
11
';
•
T
•
43
No
63
Yes
48
W
46
3900
20
E
25
19.8
16
NE
78
Y:',%-,,
48
21.4
,.-
,. -
2%. -
Mtt
2 .:
3700
,A,,
,
4200
30
NE
tx:-',1,%t
4000
%
14
N
No
_
1`07
19.2
88
No
61
No
'''',2t,,
3
19.5
66
Habitat T I
GRi1C8lif'
ABGR/LIBO/LIBO
591)
PSMENACA 250.
ABGR/LIBO (590)
SME/VACA 250
PSME/SYAL/AGSP
311
Sm5 1G'
PSME/VACA 250
:i.PSMENACA 250
PSME/VACA (250)
R/LT80,: 590):Vt
PSME/CARU 350
,,TSME/PHMA:: 260 —1
"58'2
4-'
PSME/CAGE 330
R/L
A,4,4
p,:.0.-.,
.
'BA= square feet of basal area per acre.
Series/Habitat Type/Phase (Pfister et.al. 1977).
3 Transect 4 was burned in the 1998 Boyer Wildfire and was excluded from further analyses.
(average needle complement, p<0.001; average
needle length, p<0.01). Average twig/branch
mortality significantly worsened (p<0.05) and
there was no change in needle color from 1993 to
1998. Trees put on significant diameter growth
over the 5 years; average d.b.h. increased from
21 inches to 22.4 inches (Table 3).
Average cumulative crown rating for all tagged
trees at time of transect establishment was 1.8.
When live trees were rated again 5 years later,
cumulative crown rating significantly improved
to 1.6 (p<0.001). Two of the individual crown
characteristics also significantly improved
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5
•
Yes
E
Crown Conditions
•
46
30
0
27
" 440;
15
19.8
3600
0'.i.:Z
3400
2'413
No
N
,
•
67
20
.
6
tg 9:9
23.0
A
Asiect
.,.,,
43
confirmed
on the
transect
%
;.,
H. annosum
Average
BA' for
all trees
f lg
91
Pine Trees
>=15.0"d.b.h.
i.
48
sy
•
1993
Average
d.b.h.
inches
2 5.
20.3
Table 3. Average crown characteristics for trees alive in 1993 and 1998 on randomly selected transects
(excluding Transect 4).
1993 Average Value (SE')
1998 Average Value (SE')
Needle Complement
0.37 (0.020)
0.19 (0.016)
Needle Len: h
Needle Color
0.19 (0.020)
0.02 (0.007)
0.14 0.018
0.02 (0.006)
Twig/Branch Mortality
1.20 (0.020)
1.25 (0.021)
Cumulative Crown Rating
1.79 (0.048)
1.61 (0.046)
Crown Characteristic
D.B.H.
Trend
(p value)
Improved*
(p<0.001)
Improved*
s<0.01
No Change
Worsened*
(p<0.05)
Improved*
(p<0.001)
22.42" (0.203)
21.92" (0.201)
SE = standard error of the mean
* significantly different
Darker). Almost 6% of all live trees had
Elytroderma recorded as a damaging agent.
Other Damages
Three damages were recorded for each tagged
tree. Table 4 lists the five most common agents
recorded for all transects and their frequencies.
In addition to the above damaging agents
recorded for each tree, H. annosum was
confirmed on 20% of transects.
Table 4. Most common damages recorded for
live trees at plot establishment.
i
In 1994 (year 1), one tree was killed by
secondary bark beetles and tested positive for H.
annosum. In 1995 (year 2), one tree was killed
by a combination of lightning and secondary
bark beetles, and two trees were salvaged after
death so cause of mortality could not be
confirmed, but was assumed to be bark beetles.
Samples were collected from the lightning killed
tree and the stumps of the two salvaged trees,
but H. annosum was not confirmed. No trees
died in 1996 (year 3). In 1997/1998 (years 4 and
5), one tree was killed by western pine beetles
and two trees were lost to road improvement and
harvesting. Samples were not collected from
these three trees.
Fre uenc
1993 Dama e
217
Western I an rust'
67
Fire scars
No damage
Mortality
-
311
Western gall rust = Endocronartium harknessii
(Moore) Hirat.
Elytroderma needle blight = Elytroderma deformans
(Weir) Darker
The most common biological damaging agent
recorded was western gall rust (Endocronartium
harknessii (Moore) Hirat.). Over 21% of trees
had western gall rust recorded as a damaging
agent. The second most common biological
damaging agent recorded was Elytroderma
needle blight (Elytroderma deformans (Weir)
Transect 4 was burned by the Boyer Wildfire
during the 1998 fire season. Seven trees on the
transect were killed outright by the fire.
Although many trees survived, the transect was
6
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improvement; and five trees died from "natural"
causes other than fire. The latter five trees were
the only trees we could use to calculate a
representative mortality rate. Average annual
mortality was 0.14% for ponderosa pine trees
15.0 inches d.b.h. or greater. See Table 5 for a
summary of mortality.
no longer useful for this project, so the 1998
remeasurement for transect 4 was dropped from
the analysis.
Over the 5-year period, 14 trees died from
various causes: seven trees were lost to fire; two
trees were lost to harvesting and/or road
Table 5.
Individual mortality characteristics and probable cause .
Year of
Mortali
1993 D.B.H.
inches
Tree ID'
1
1994
1993
Cumulative
Crown Ratin .
Presence of
H. annosum
Cause
Beetles
Salva e
Li htnin /beetles
-
g
g
g
*-4
4
,
.,
.
,
.,,
.
,
-
,
,
IN.
97/9 g.,
1997/98
1
,
.,
10-426
-.4
5
22.4
r.,:,,,
',.
"1-3997.1.9
V_
i.
, .
:Eit-0,'
.
,
0
4
.
,
, ,
tt
..
.
,
.
,
,
.
,
'
vca
oZt
Not sam led
Beetles
s
awe
ii, •
e
161.
.rovem.,
I Tree ID=Transect number - tree number.
2 Mortality in shaded rows were not included in calculating mortality rate nor decline symptoms tied to
mortality.
(Table 6). Additionally, trees that died were
significantly larger in d.b.h. than trees that
survived (26.9 inches and 21.92 inches,
respectively; p<0.05).
When 1993 average crown characteristics of
trees that died over 5 years were compared to
trees that lived over 5 years, they were found to
be significantly different. All crown ratings were
statistically greater (worse) for trees that died
Table 6.
1993 average crown characteristics for trees dead through 1998 and 1998 live (excluding
Transect 4).
Crown Characteristic
Needle Complement
Needle Length
Needle Color
Twig/Branch Mortality
Cumulative Crown Rating
D.B.H.
Dead Tree Average
Value (SE')
1.20 (0.490)
0.80 (0.374)
0.60 (0.400)
2.20 (0.490)
4.80 (1.463)
26.9" (2.277)
SE - standard error of the mean
7
Live Tree Average
Value (SE')
0.37 (0.020)
0.19 (0.020)
0.02 (0.007)
1.20 (0.020)
1.79 (0.048)
21.92" (0.201)
Significance
p<0.001
p=0.01
P<0.001
p<0.001
p<0.001
p<0.05
•
consistently worsened over 5 years, although
only three measurements were statistically
significant: needle color, twig/branch mortality
and cumulative crown rating (Table 8). The
average d.b.h. increased from 18.9 inches to
19.9 inches over the 5 years.
Annosum Transect
This transect was selected by the Tribes based
on the presence of H. annosum. Table 7 lists the
transect characteristics.
•
The non-randomly selected Annosum transect
was analyzed separately, but similarly to other
transects. Average crown characteristics
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Table 7. Annosum transect characteristics at time of plot establishment.
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#Ponderosa
Pine Trees
Elevation Slope
(feet)
(%) As ect >=15.0"d.b.h.
255
3200
10
SE
1993
Ave.
d.b.h.
(inches)
18.9
H.annosum
Average
BA' for
all .ees
100
confirmed
on the
transect
Yes
Habitat T Ee2
PSME/SYAL
(310)
' BA= square feet of basal area per acre.
2 Series/Habitat Type/Phase (Pfister et.al. 1977).
•
Table 8. Average tree characteristics for trees alive in 1993 and 1998 on Annosum Transect only.
1993 Average Value (SE')
1998 Average Value (SE')
Needle Complement
0.11 (0.020)
0.12 (0.024)
Needle Length
0.04 (0.013)
0.06 (0.017)
Needle Color
0.01 (0.006)
0.05 (0.015)
Twig/Branch Mortality
1.06 (0.018)
1.20 (0.030)
Cumulative Crown Rating
1.22 (0.037)
1.43 (0.065)
18.86" (0.216)
19.90" (0.323)
Crown Characteristic
•
D.B.H.
SE = standard error of the mean
2 NS = not significant at p<= 0.10
* significantly different
8
Trend
(p value')
Worsened
(NS)
Worsened
(NS)
Worsened*
(p=0.01)
Worsened*
(p<0.001)
Worsened*
((p----0.001)
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Only one tree died over 5 years, and it occurred
in 1994 (Table 9). The tree had been hit by
lightning at time of plot establishment and was
attacked by western pine beetles (Dendroctonus
brevicomis LeConte) the following year. In
1994, we collected a 2-inch thick disc at ground
level which tested positive for S. meineckellus.
Death of one tree on this transect equates to an
average annual mortality rate of 0.08%. Because
only one tree died, a comparison of crown
characteristics of trees that lived versus trees that
died was not made.
When the non-randomly selected Annosum
transect was compared to the 14 randomly
selected transects, trees on the Annosum transect
were significantly smaller in d.b.h. and average
crown characteristics were significantly better
than trees on the other transects (Table 10).
Table 9. Individual mortality characteristics and probable cause .
Year of
Mortality
1994'
Tree ED'
SS-163
1993 D.B.H.
(inches)
20.8
1993
Cumulative
Crown Rating
1
•
•
Cause
Lightning/beetles
H. annosum
Yes
Tree ID=Transect number - tree number. (SS=Annosum Transect)
Table 10. 1993 average live tree characteristics for the non-randomly selected Annosum Transect
compared to the 14 randomly selected transects at plot establishment.
Crown Characteristic
Needle Complement
Needle Length
Needle Color
Twig/Branch Mortality
Cumulative Crown
Rating
D.B.H.
1993 Average Value (SE')
Annosum Transect
0.11 (0.020)
0.04 (0.013)
0.01 (0.006)
1.06 (0.018)
1993 Average Value (SE)
14 Randomly Selected
Transects
0.37 (0.020)
0.20 (0.020)
0.03 (0.007)
1.21 (0.021)
Significance
p<0.001
p<0.001
p<0.10
p<0.001
1.22 (0.036)
1.81 (0.049)
p<0.001
18.86" (0.216)
21.96" (0.201)
p<0.001
SE = standard error of the mean
DISCUSSION
•
Presence of
from these transects indicate the mortality rate
was low for the 5 years of this study (19931998). Mortality and decline rates may have
been higher in previous years, and the period of
this study did not capture those higher rates.
Overall, there was very little decline or mortality
in ponderosa pine trees 15 inches or greater
d.b.h. over the 5 years of this study. When all
randomly selected transects were analyzed,
crown characteristics generally improved. The
exception was the characteristic of twig and
branch mortality, which worsened. The
appearance of a decline noted in the late 1980's
and early 1990's was likely very real, but data
The appearance of an overall decline in the
previous years may have been due to droughty
conditions the area experienced in the late
1980's and early 1990's. Data from the National
Climatic Data Center indicates a droughty
9
damaging in trees of low vigor with poor
crowns, however, the disease is often more
conspicuous in good crowns. In larger trees,
direct mortality from Elytroderma is uncommon,
but infection contributes to permanent
deterioration of crowns and consequent
reduction of growth and increase in vulnerability
to bark beetles, root diseases and other enemies
(Childs 1968; Childs et al. 1971). There is also
some evidence to suggest that very droughty
conditions, such as occurred in the area prior to
plot establishment, hasten death of infected
twigs (Childs 1968).
period existed from 1986 through the second
half of 1993, the year of plot establishment. At
plot establishment in 1993, 3 of the previous 6
years experienced moderate drought and the
remaining 3 years were barely within normal
range, according to the Palmer Drought Severity
Index (National Climatic Data Center 2000).
Palmer Drought Severity Index ranged from –
5.0 (severe drought) during 1987, to +1.0 (near
normal) during 1989-91. During the 5-year time
period of this study, precipitation for the area
had returned to more normal levels, with 2 of 5
years falling in a "very moist spell" (Palmer
Drought Severity Index = +3.0 to +3.9, or
greater). Another period of drought in the future
may initiate another appearance of decline.
The addition of the annosum transect allowed
some comparisons to be made between a known
annosum-infected site with randomly selected
sites. On the annosum transect, average crown
characteristics consistently worsened from 1993
to 1998. This is contrary to randomly selected
transects, where average crown characteristics
consistently improved over the 5 years. It is
possible that if we analyzed the randomly
selected transects with confirmed annosum
infection separately, we might see a similar
trend to the non-randomly selected annosum
transect. This warrants further investigation.
Crowns of trees that died were significantly
worse than trees that survived the 5-year time
period. This indicates the rating system used in
this study may have some predictive value.
Individual crown characterisitics which are most
predictive cannot be determined from this study
due to the low mortality rate. A most predictive
characteristic and threshold may be revealed
over time.
Twig and branch mortality worsened over the
study period. This general twig and branch
decline may be due to varying levels of western
gall rust and Elytroderma needle blight. Western
gall rust was recorded on over 21% of all live
trees in 1993, and Elytroderma needle blight was
recorded on almost 6% of the trees. Western gall
rust causes galls to form on infected branches or
stems of trees. Branches and sometimes small
stems are killed when insects and other fungi
attack galled tissue (Byler et al. 1972). High
levels of western gall rust in crowns of trees can
cause significant branch mortality and possibly
top kill (Peterson et al. 1960; Hennon 1993),
contributing to an overall decline in the vigor of
individual trees.
The 1993 crown measurements for the annosum
transect were statistically better than the
randomly selected transects. The trees on the
annosum transect are smaller and appear to be
younger (data not analyzed), so they have not
been exposed to damaging agents for as long as
trees on the other transects. It is noteworthy that
trees on the annosum transect have healthier
crowns than trees on the randomly selected
transects, but showed a general crown decline
over 5 years, rather than an improvement in
crown conditions as found on the randomly
selected transects.
FUTURE PLANS
We plan to monitor these transects for another 5year interval, at which time their usefulness will
be assessed.
Elytroderma needle blight is considered the most
important needle disease of pines in the Pacific
Northwest (Childs et al. 1971). It is most
10
•
small pores. "Button conks" are small, corky
mounds of sterile tissue, and are generally cream
colored (Hagle et al. 1987).
ACKNOWLEDGEMENTS
•
•
•
•
We acknowledge Sue Hagle for initiating the
project and providing her expertise in project
planning. We also acknowledge the
Confederated Salish and Kootenai Tribes for
assisting in the installation of transects. We
extend our thanks to Terry Reedy for his
assistance in project management and field data
collection, Ka Moua for keying data into our
data management system, and Michael Marsden,
INTECS International, Inc. employee under
contract with Forest Health Technology
Enterprise Team, for performing the statistical
consulting and analyses. We acknowledge many
others who were involved in field data
collection, including: Jane Taylor, Ken Gibson,
Bill Cramer, Carol Randall, Tim McConnell,
and various seasonal workers.
Annosum decay varies, but is generally
recognized in its early stage as a stain in the
outer heartwood, which can be readily seen in a
cross section of a freshly cut stump. The
advanced decay is often white to yellow, stringy
to laminate, and may contain white pockets with
black flecks (Schmitt et al. 1984, Hagle et al.
1987). Positive identification of the decay can
be made by culturing and/or incubating decayed
wood and observing the imperfect stage of the
fungus, Spiniger meineckellus (A.J. Olson)
Stalpers.
H annosum
in western North America consists
of two intersterility groups or biological species.
These two types, 's' and 'p', have very different
host specificities. Hosts for the p-type include
pine species, incense cedar, and western juniper.
The hosts for the s-type include true firs, giant
sequoia, spruce, western hemlock, and Douglasfir. This host specificity is not apparent in
stump infections; both groups have been isolated
from "non-host" stumps without causing disease
in neighboring "host" trees (Lockman 1993,
Otrosina et al. 1992, Klielunas 1986). In
intermountain forests, butt rot is associated with
infection in spruce and true firs, and outright tree
mortality is more common in pines and Douglasfir (Hagle 1985, Byler and Hagle 1985).
Infected trees often are attacked by bark beetles
(Byler 1989, Ferrell and Parmeter 1989,
Hadfield et al. 1986).
APPENDIX
•
•
•
•
•
•
Identification and Biology of Annosum Root
Disease
Annosum root disease is a white root and butt
rot of many conifer species throughout western
United States and many other temperate forest
ecosystems (Hodges 1969). It is caused by the
fungus Heterobasidion annosum (Fr.)Bref..
Symptoms of annosum root disease are similar
to other root diseases. Infected trees may have
faded, chlorotic crowns, reduced growth, stressinduced cone crops, resin at the base, resinsoaked and discolored wood, and decay in roots
and butts. Annosum root disease can be
identified by its fruiting bodies, characteristic
decay it causes, and by isolating H annosum
from the associated incipient decay.
Annosum root disease can spread in several
ways. H. annosum in a diseased tree can infect a
healthy neighboring tree by ectomycelium
growing across root contacts, if the neighboring
tree is a susceptible species. H annosum can
also spread by airborne spores (Hodges 1969,
Hsiang et al. 1989). When spores land on a
newly cut stump or fresh basal wound, they may
germinate and colonize the wood if conditions
are favorable. Spore infections are usually
limited to the tops of freshly cut stumps in pine
H annosum
fruiting bodies (i.e., sporophores or
conks) can sometimes be found inside decayed
hollowed stumps, just under the duff layer at the
base of infected pines. It may also be found as
small "button conks" on the outside of infected
roots and root collars on all susceptible species.
Annosum fruiting bodies are perennial, woody
to leathery, with dark brown upper surface and
white to cream-colored lower surface with very
11
Ferrell, G.T. and J.R. Parmeter, Jr. 1989.
Interactions of root disease and bark beetles.
pp. 105-108. /N: Proceedings on the
Symposium on Research and Management of
Annosus Root Disease (H. annosum) in
western North America. Monterey, California.
April 18-21, 1989. Gen. Tech. Rep. PSW-116.
species (Otrosina and Cobb 1989), while freshly
cut stumps and fresh basal wounds are
susceptible to infection in other species (Schmitt
et al. 1984). The newly infected stump or tree
may then initiate a new disease center by rootto-root contact.
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13